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Richard C. Brusca, PhD
Executive Director Emeritus, Arizona-Sonora Desert Museum and Research Scientist, Department of Ecology and Evolutionary Biology, University of Arizona
Gonzalo Giribet, PhD
Director, Museum of Comparative Zoology and Alexander Agassiz Professor of Zoology, Harvard University
Wendy Moore, PhD
Associate Professor and Curator, Department of Entomology, University of Arizona with illustrations by Nancy Haver
About the cover
Day Octopus (Octopus cyanea).
Found throughout the tropical Indo-Pacific, this is the most commonly seen octopus in Hawai’i. Hidden in a lair on the reef floor by night, the octopus will reach out with tentacles to gather coral rubble to protect the entrance. When at rest in its lair, it is a uniformly dark reddish-brown but during the day when hunting crabs on the reef bottom, it displays a spectacular array of color patterns and textures to match the varied landscape it crosses. An ink cloud released during escape contains mucus, melanin, dopamine, and tyrosinase, and is thought to confuse a fish predator and to depress its olfactory ability.
Larry Jon Friesen, PhD, completed his graduate research at the University of California, Santa Barbara, in Animal Communication. Dr. Friesen is a Professor of Biological Sciences at Santa Barbara City College, teach ing Natural History, Evolution and Animal Diversity and continuing his life-long passion for nature photography.
Invertebrates, Fourth Edition
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Library of Congress Cataloging-in-Publication Data
Names: Brusca, Richard C., author. | Giribet, Gonzalo, author. | Moore, Wendy, author. Title: Invertebrates / Richard C. Brusca, PhD, Executive Director Emeritus, ArizonaSonora Desert Museum, Research Scientist, Department of Ecology and Evolutionary Biology, University of Arizona, Gonzalo Giribet, PhD, Director, Museum of Comparative Zoology and Alexander Agassiz Professor of Zoology, Harvard University, Wendy Moore, PhD, Associate Professor and Curator, Department of Entomology, University of Arizona.
Description: Fourth edition. | New York : Oxford University Press, [2023] | Revised edition of: Invertebrates / Richard C. Brusca, Wendy Moore, Stephen M. Shuster. Third edition. 2016. | Includes bibliographical references and index.
Identifiers: LCCN 2021054094 | ISBN 9780197554418 (hardback) | ISBN 9780197637173 (epub)
Subjects: LCSH: Invertebrates.
Classification: LCC QL362 .B924 2023 | DDC 592--dc23/eng/20211103
LC record available at https://lccn.loc.gov/2021054094
9 8 7 6 5 4 3 2 1
Printed in the United States of America
for more ebook/ testbank/ solution manuals requests: email 960126734@qq.com
We dedicate this book to all our fellow teachers and students of invertebrate zoology around the world.
Brief Contents
CHAPTER 1 Introduction 1
CHAPTER 2 Systematics, Phylogeny, and Classifications 27
CHAPTER 3 Introduction to the Animal Kingdom: Animal Architecture and Body Plans 43
CHAPTER 4 Introduction to the Animal Kingdom: Development, Life Histories, and Origin 91
CHAPTER 5 Phylum Porifera: The Sponges 119
CHAPTER 6 Two Enigmatic Phyla: Placozoa and Ctenophora (The Comb Jellies) 165
CHAPTER 7 Phylum Cnidaria: Anemones, Corals, Jellyfish, and Their Kin 185
CHAPTER 8 A Brief Introduction to the Bilateria and Its Major Clades 245
CHAPTER 9 Phylum Xenacoelomorpha: Basal Bilaterians 249
CHAPTER 10 Protostomia, Spiralia, and the Phylum Dicyemida 273
CHAPTER 11 Gnathifera: The Phyla Gnathostomulida, Rotifera (including Acanthocephala), Micrognathozoa, and Chaetognatha 281
CHAPTER 12 Platytrochozoa and Two Enigmatic Phyla: Entoprocta and Cycliophora 311
CHAPTER 13 Introduction to the Lophotrochozoa, and the Phylum Mollusca 321
CHAPTER 14 Phylum Nemertea: The Ribbon Worms 397
CHAPTER 15 Phylum Annelida: The Segmented (and Some Unsegmented) Worms 415
CHAPTER 16 The Lophophorates: Phyla Phoronida, Bryozoa, and Brachiopoda 487
CHAPTER 17 Rouphozoa: The Phyla Platyhelminthes (Flatworms) and Gastrotricha (Hairy-Bellied Worms) 519
CHAPTER 18 Introduction to Ecdysozoa: Scalidophora (Phyla Kinorhyncha, Priapula, Loricifera) 563
CHAPTER 19 Nematoida: Phyla Nematoda and Nematomorpha 579
CHAPTER 20 Panarthropoda and the Emergence of the Arthropods: Tardigrades, Onychophorans, and the Arthropod Body Plan 607
CHAPTER 21 Phylum Arthropoda—Subphylum Crustacea: Crabs, Shrimps, and Their Kin 659
CHAPTER 22 Phylum Arthropoda—Subphylum Hexapoda: Insects and Their Kin 735
CHAPTER 23 Phylum Arthropoda—Subphylum Myriapoda: Centipedes, Millipedes, and Their Kin 785
CHAPTER 24 Phylum Arthropoda: Subphylum Chelicerata 801
CHAPTER 25 Introduction to Deuterostomia, and the Phylum Hemichordata 857
CHAPTER 26 Phylum Echinodermata : Starfish, Sea Urchins, Sea Cucumbers, and their Kin 873
CHAPTER 27 Phylum Chordata: Cephalochordata and Urochordata 911
CHAPTER 28 Perspectives on Invertebrate Phylogeny 935
Classification of the Animal Kingdom (Metazoa) xx
A Phylogeny of Metazoa xxi
Geologic Time Scale xxii
CHAPTER 1 Introduction 1
Keeping Track of Life 3
Prokaryotes and Eukaryotes 7
Where Did Invertebrates Come From? 9
The Dawn of Life 10
The Ediacaran Period and the Origin of Animals 10
The Paleozoic Era (541–251.9 Ma) 11
The Mesozoic Era (251.9–66 Ma) 15
The Cenozoic Era (66 Ma–present) 16
Where Do Invertebrates Live? 16
Marine Habitats 16
Estuaries and Coastal Wetlands 21
Freshwater Habitats 21
Terrestrial Habitats 22
A Special Type of Environment: Symbiosis 22
Changing Views of Invertebrate Phylogeny 24
Legacy Names 25
Phylogenetics and Classification Schemes 25
A Final Introductory Message to the Reader 25
CHAPTER 2 Systematics, Phylogeny, and Classifications 27
Phylogeny, Monophyly, Paraphyly, and Polyphyly 28
Homology 29
Apomorphy and Plesiomorphy 32
Challenges of Phylogenetic Inference 32
Constructing Phylogenies 33
Biological Classification 35
Nomenclature 38
CHAPTER 3 Introduction to the Animal Kingdom: Animal Architecture and Body Plans 43
Body Symmetry 44
Cellularity, Body Size, Germ Layers, and Body Cavities 47
Locomotion and Support 49
Reynolds Number 49
Ameboid Locomotion 50
Cilia and Flagella 50
Muscles and Skeletons 52
Feeding and Digestion 56
Intracellular and Extracellular Digestion 56
Feeding Strategies 57
Excretion and Osmoregulation 65
Nitrogenous Wastes and Water Conservation 66
Osmoregulation and Habitat 66
Excretory and Osmoregulatory Structures 67
Circulation and Gas Exchange 69
Internal Transport 69
Circulatory Systems 70
Hearts and Other Pumping Mechanisms 71
Gas Exchange and Transport 71
Nervous Systems and Sense Organs 75
Sense Organs 76
Independent Effectors 81
Bioluminescence 81
Nervous Systems and Body Plans 81
Hormones and Pheromones 84
Reproduction 84
Asexual Reproduction 84
Sexual Reproduction 86
Parthenogenesis 88
CHAPTER 4
Introduction to the Animal Kingdom: Development, Life Histories, and Origin 91
Evolutionary Developmental Biology: Evo-Devo 92
Developmental Tool Kits 92
The Relationship Between Genotype and Phenotype 93
The Evolution of Novel Gene Function 93
Gene Regulatory Networks 93
Eggs and Embryos 95
Eggs 95
Cleavage 95
Orientation of Cleavage Planes 96
Radial and Spiral Cleavage 96
Cell Fates 99
Blastula Types 101
Gastrulation and Germ Layer Formation 101
Mesoderm and Body Cavities 103
Life Cycles: Sequences and Strategies 105
Classification of Life Cycles 105
Indirect Development 107
Settling and Metamorphosis 107
Direct Development 108
Mixed Development 108
Adaptations to Land and Fresh Water 109
Parasite Life Cycles 109
The Relationships Between Ontogeny and Phylogeny 110
The Concept of Recapitulation 110
Heterochrony and Paedomorphosis 111
The Origin of the Metazoa 112
Origin of the Metazoan Condition 112
Historical Perspectives on Metazoan Origins 112
The Origin of Multicellularity 114
The Origin of the Bilateral Condition and the Coelom 115
The Trochaea Theory 116
Closing Thoughts 117
CHAPTER 5 Phylum Porifera: The Sponges 119
Phylum Porifera: The Sponges 120
Taxonomic History and Classification 123
The Poriferan Body Plan 126
Body Structure and the Aquiferous System 127
More on Sponge Cell Types 132
Support 136
Nutrition, Excretion, and Gas Exchange 138
Activity and Sensitivity 143
Reproduction and Development 143
Some Additional Aspects of Sponge Biology 154
Distribution and Ecology 154
Biochemical Agents 154
Growth Rates 155
Symbioses 156
Poriferan Phylogeny 159
The Origin of Sponges 159
Evolution within the Porifera 160
CHAPTER 6 Two Enigmatic Phyla: Placozoa and Ctenophora (The Comb Jellies) 165
Phylum Placozoa 166
Phylum Ctenophora 167
Taxonomic History and Classification 169
The Ctenophoran Body Plan 172
Support and Locomotion 175
Feeding and Digestion 176
Circulation, Excretion, Gas Exchange, and Osmoregulation 179
Nervous System and Sense Organs 179
Reproduction and Development 181
Ctenophoran Phylogeny 183
CHAPTER 7 Phylum Cnidaria: Anemones, Corals, Jellyfish, and Their Kin 185
Taxonomic History and Classification 190
The Cnidarian Body Plan 196
The Body Wall 197
Support 209
Movement 212
Cnidae 215
Feeding and Digestion 218
Defense, Interactions, and Symbiosis 220
Circulation, Gas Exchange, Excretion, and Osmoregulation 227
Nervous System and Sense Organs 227
Reproduction and Development 231
Cnidarian Evolutionary History 240
Earliest Cnidaria 240
Cnidarian Phylogeny 241
CHAPTER 8 A Brief Introduction to the Bilateria and Its Major Clades 245
The Bilateria 245
Deuterostomes and Protostomes 246
CHAPTER 9 Phylum Xenacoelomorpha: Basal Bilaterians 249
The Basal Bilaterian 249
Phylum Xenacoelomorpha 250
Subphylum Acoelomorpha 252
Class Acoela 252
The Acoel Body Plan 255
Body Wall and External Appearance 255
Body Musculature, Support, and Movement 256
Nutrition, Excretion, and Gas Exchange 257
Nervous Systems and Sense Organs 258
Reproduction and Development 259
Class Nemertodermatida 261
The Nemertodermatid Body Plan 263
Body Structure 263
Cell and Tissue Organization 263
Support and Movement 263
Nutrition, Excretion, Gas Exchange 264
Nervous System 265
Reproduction and Development 265
Subphylum Xenoturbellida 267
The Xenoturbellid Body Plan 268
General Body Structure 268
Support and Movement 269
Nutrition, Excretion, and Gas Exchange 270
Nervous System and Sense Organs 270
Reproduction and Development 270
CHAPTER 10 Protostomia, Spiralia, and the Phylum Dicyemida 273
Protostomes and Deuterostomes 273
Spiralia and Ecdysozoa 274
The Phylum Dicyemida (= Rhombozoa) 275
Anatomy and Biology of Dicyemidans 275 Life Cycles 277
CHAPTER 11 Gnathifera: The Phyla Gnathostomulida, Rotifera (including Acanthocephala), Micrognathozoa, and Chaetognatha 281
Phylum Gnathostomulida: The Gnathostomulids 283
The Gnathostomulid Body Plan 284
Body Wall, Support, and Locomotion 284
Nutrition, Circulation, Excretion, and Gas Exchange 284
Nervous System 284
Reproduction and Development 284
Phylum Rotifera: The Free-Living Rotifers 284
The Rotifer Body Plan 286
Body Wall, General External Anatomy, and the Corona 286
Body Cavity, Support, and Locomotion 287
Feeding and Digestion 288
Circulation, Gas Exchange, Excretion, and Osmoregulation 289
Nervous System and Sense Organs 290
Reproduction and Development 290
Phylum Rotifera, Subclass Acanthocephala: The Acanthocephalans 292
The Acanthocephalan Body Plan 293
Body Wall, Support, Attachment, and Nutrition 293
Circulation, Gas Exchange, and Excretion 294
Nervous System 294
Reproduction and Development 294
Phylum Micrognathozoa: The Micrognathozoans 295
The Micrognathozoan Body Plan 296
Epidermis, Ciliation, and Body Wall Musculature 296
Locomotion 298
Pharyngeal Apparatus, Feeding, and Digestion 298
Circulation, Gas Exchange, and Excretion 298
Nervous System and Sense Organs 301
Reproduction and Development 301
Phylum Chaetognatha 301
The Chaetognath Body Plan 304
Body Wall, Support, and Movement 304
Feeding and Digestion 306
Circulation, Gas Exchange, and Excretion 306
Nervous System and Sense Organs 306
Reproduction and Development 307
CHAPTER 12 Platytrochozoa and Two Enigmatic Phyla: Entoprocta and Cycliophora 311
Phylum Entoprocta: The Entoprocts 312
The Entoproct Body Plan 314
Body Wall, Support, and Movement 314
Feeding and Digestion 314
Circulation, Gas Exchange, and Excretion 314
Nervous System 315
Reproduction and Development 316
Phylum Cycliophora: The Cycliophorans 317
CHAPTER 13 Introduction to the Lophotrochozoa, and the Phylum Mollusca 321
Phylum Mollusca 322
Taxonomic History and Classification 322
The Molluscan Body Plan 344
The Body Wall 346
The Mantle and Mantle Cavity 346
The Molluscan Shell 347
Torsion, or “How the Gastropod Got its Twist” 353
Locomotion 356
Feeding 361
Digestion 370
Circulation and Gas Exchange 373
Excretion and Osmoregulation 377
Nervous System 378
Sense Organs 380
Cephalopod Coloration and Ink 384
Reproduction 385
Development 389
Molluscan Evolution and Phylogeny 392
CHAPTER 14 Phylum Nemertea: The Ribbon Worms 397
Taxonomic History and Classification 399
Classification 399
The Nemertean Body Plan 400
Body Wall 401
Support and Locomotion 402
Feeding and Digestion 402
Circulation and Gas Exchange 406
Excretion and Osmoregulation 406
Nervous System and Sense Organs 408
Reproduction and Development 409
Nemertean Phylogeny 411
CHAPTER 15 Phylum Annelida: The Segmented (and Some Unsegmented) Worms
Taxonomic History and Classification 416
The Annelid Body Plan 426
Body Forms 426
Body Wall and Coelomic Arrangement 428
Support and Locomotion 429
Feeding and Digestion 432
Circulation and Gas Exchange 441
Excretion and Osmoregulation 444
Nervous System and Sense Organs 446
Reproduction and Development 450
Sipuncula: The Peanut Worms 457
Classification of Sipuncula 459
The Sipunculan Body Plan 460
Body Wall, Coelom, Circulation, and Gas Exchange 460
Support and Locomotion 461
Feeding and Digestion 462
Excretion and Osmoregulation 462
Nervous System and Sense Organs 463
Reproduction and Development 463
415
Thalassematidae: The Spoon Worms 465
Body Wall and Coelom 465
Support and Locomotion 465
Feeding and Digestion 465
Circulation and Gas Exchange 469
Excretion and Osmoregulation 469
Nervous System and Sense Organs 469
Reproduction and Development 469
Siboglinidae: Vent Worms and Their Kin 470
Siboglinid Taxonomic History 473
The Siboglinid Body Plan 473
The Tube, Body Wall, and Body Cavity 473
Nutrition 474
Circulation, Gas Exchange, Excretion, and Osmoregulation 474
Nervous System and Sense Organs 474
Reproduction and Development 474
Hirudinea: Leeches and Their Relatives 476
The Hirudinean Body Plan 477
Body Wall and Coelom 477
Support and Locomotion 477
Feeding and Digestion 478
Circulation and Gas Exchange 479
Excretion and Osmoregulation 480
Nervous System and Sense Organs 480
Reproduction and Development 481
Orthonectida: Extremely Simplified Annelids 482
Annelid Phylogeny 483
CHAPTER 16 The Lophophorates: Phyla Phoronida, Bryozoa, and Brachiopoda 487
Taxonomic History of the Lophophorates 488
The Lophophorate Body Plan 489
Phylum Phoronida: The Phoronids 490
The Phoronid Body Plan 490
Body Wall, Body Cavity, and Support 490
The Lophophore, Feeding, and Digestion 494
Circulation, Gas Exchange, and Excretion 494
Nervous System 495
Reproduction and Development 495
Phylum Bryozoa: The Moss Animals 496
The Bryozoan Body Plan 499
The Body Wall, Coelom, Muscles, and Movement 501
Zooid Interconnections 502
The Tentacle Crown, Feeding, and Digestion 503
Circulation, Gas Exchange, and Excretion 504
Nervous System and Sense Organs 505
Reproduction and Development 506
Phylum Brachiopoda: The Lamp Shells 509
The Brachiopod Body Plan 512
The Body Wall, Coelom, and Support 512
The Lophophore, Feeding, and Digestion 513
Circulation, Gas Exchange, and Excretion 514
Nervous System and Sense Organs 515
Reproduction and Development 515
CHAPTER 17 Rouphozoa: The Phyla Platyhelminthes (Flatworms) and Gastrotricha (Hairy-Bellied Worms) 519
Introduction to Rouphozoa 519
The Phylum Platyhelminthes (Flatworms) 520
Taxonomic History and Classification 522
The Platyhelminth Body Plan 527
Body Wall 529
Support, Locomotion, and Attachment 532
Feeding and Digestion 533
Circulation and Gas Exchange 537
Excretion and Osmoregulation 538
Nervous System and Sense Organs 539
Reproduction and Development 541
Platyhelminth Phylogeny 554
Phylum Gastrotricha: The Gastrotrichs, or HairyBellied Worms 556
The Gastrotrich Body Plan 558
Body Wall 558
Support and Locomotion 558
Feeding and Digestion 558
Circulation, Gas Exchange, Excretion, and Osmoregulation 558
Nervous System and Sense Organs 558
Reproduction and Development 560
CHAPTER 18 Introduction to Ecdysozoa: Scalidophora (Phyla Kinorhyncha, Priapula, Loricifera) 563
Introduction to Ecdysozoa 563
The Scalidophora 564
Phylum Kinorhyncha: The Kinorhynchs, or Mud Dragons 564
The Kinorhynch Body Plan 567
Body Wall 567
Support and Locomotion 567
Feeding and Digestion 567
Circulation, Gas Exchange, Excretion, and Osmoregulation 567
Nervous System and Sense Organs 568
Reproduction and Development 568
Phylum Priapula: The Priapulans, or Penis Worms 568
Priapulan Body Plan 570
Body Wall, Support, and Locomotion 570
Feeding and Digestion 571
Circulation, Gas Exchange, Excretion, and Osmoregulation 571
Nervous System and Sense Organs 572
Reproduction and Development 572
Phylum Loricifera: The Loriciferans 572
CHAPTER 19 Nematoida: Phyla Nematoda and Nematomorpha 579
Phylum Nematoda: Roundworms 581
Classification of Phylum Nematoda 582
The Nematode Body Plan 586
Body Wall, Support, and Locomotion 586
Feeding and Digestion 588
Circulation, Gas Exchange, Excretion, and Osmoregulation 590
Nervous System and Sense Organs 592
Reproduction, Development, and Life Cycles 594
Life Cycles of Some Parasitic Nematodes 597
CHAPTER
20 Panarthropoda and
Phylum Nematomorpha: Horsehair Worms and Their Kin 600
The Nematomorph Body Plan 601
Body Wall, Support, and Locomotion 601
Feeding and Digestion 603
Circulation, Gas Exchange, Excretion, and Osmoregulation 603
Nervous System and Sense Organs 604
Reproduction and Development 604
the Emergence of the Arthropods: Tardigrades, Onychophorans, and the Arthropod Body Plan 607
Phylum Tardigrada 610
The Tardigrade Body Plan 613
Locomotion 615
Feeding, Digestion, and Excretion 616
Circulation and Gas Exchange 616
Nervous System and Sense Organs 616
Reproduction and Development 617
Phylum Onychophora 619
The Onychophoran Body Plan 622
Locomotion 623
Feeding and Digestion 624
Circulation and Gas Exchange 624
Excretion and Osmoregulation 625
Nervous System, Sense Organs, and Behavior 625
Reproduction and Development 626
Systematics and Biogeography 628
An Introduction to the Phylum Arthropoda 628
Taxonomic History and Classification 629
The Arthropod Body Plan and Arthropodization 630
The Body Wall 632
Arthropod Appendages 634
Support and Locomotion 636
Growth 639
The Digestive System 642
Circulation and Gas Exchange 644
Excretion and Osmoregulation 646
Nervous System and Sense Organs 647
Reproduction and Development 651
The Evolution of Arthropods 652
The Origin of Arthropods 652
Evolution within the Arthropoda 652
CHAPTER 21 Phylum Arthropoda—Subphylum Crustacea: Crabs, Shrimps, and Their Kin 659
Classification of the Crustacea 663
Synopses of Crustacean Taxa 666
The Crustacean Body Plan 699
Locomotion 703
Feeding 708
Digestive System 714
Circulation and Gas Exchange 717
Excretion and Osmoregulation 719
Nervous System and Sense Organs 720
Reproduction and Development 724
Crustacean Phylogeny 730
CHAPTER 22 Phylum Arthropoda—Subphylum Hexapoda: Insects and Their Kin 735
Classification of the Subphylum Hexapoda 738
Synopses of Hexapod Groups 739
The Hexapod Body Plan 751
General Morphology 751
Locomotion 758
The Origin of Insect Flight 761
Feeding and Digestion 762
Circulation and Gas Exchange 767
Excretion and Osmoregulation 770
Nervous System and Sense Organs 771
Reproduction and Development 775
Hexapod Evolution 780
CHAPTER 23 Phylum Arthropoda—Subphylum Myriapoda: Centipedes, Millipedes, and Their Kin 785
Myriapod Classification 787
The Myriapod Body Plan 789
Head and Mouth Appendages 791
Locomotion 791
Feeding and Digestion 791
CHAPTER
Circulation and Gas Exchange 793
Excretion and Osmoregulation 794
Nervous System and Sense Organs 794
Reproduction and Development 795
Embryonic Development 798
Myriapod Phylogeny 798
24 Phylum Arthropoda: Subphylum Chelicerata 801
Synopses of Living Chelicerate Groups 807
The Euchelicerate Body Plan 818
Spinnerets, Spider Silk, and Spider Webs 819
Locomotion 823
Feeding and Digestion 826
Circulation and Gas Exchange 831
Excretion and Osmoregulation 834
Nervous System and Sense Organs 834
Reproduction and Development 837
The Class Pycnogonida 846
The Pycnogonid Body Plan 849
External Anatomy 849
Locomotion 850
Feeding and Digestion 850
Circulation, Gas Exchange, and Excretion 852
Nervous System and Sense Organs 852
Reproduction and Development 852
Chelicerate Phylogeny 854
CHAPTER 25 Introduction to Deuterostomia, and the Phylum Hemichordata 857
Introduction to the Deuterostomia 857
Phylum Hemichordata: Acorn Worms and Pterobranchs 859
The Hemichordate Body Plan 862
Class Enteropneusta (Acorn Worms) 863
External Anatomy 863
Support Structures 863
Coelomic Cavities 863
Musculature and Locomotion 865
Feeding and Digestion 865
Circulatory System 866
Excretory System 866
Gas Exchange 866
Nervous System 866
Reproduction and Development 866
Class Pterobranchia (Pterobranchs) 868
Body Wall and Cavities 869
Support, Muscles, and Movement 869
Gut and Feeding 869
Circulation and Gas Exchange 869
Nervous System 870
Reproduction and Development 870
Hemichordate Fossil Record and Phylogeny 870
CHAPTER 26 Phylum Echinodermata: Starfish, Sea Urchins, Sea Cucumbers, and their Kin 873
Taxonomic History and Classification 877
The Echinoderm Body Plan 881
Developmental Roots of the Echinoderm Body Plan 881
Body Wall and Coelom 883
Mutable Collagenous Tissue 885
Water Vascular System 885
Support and Locomotion 887
Feeding and Digestion 889
Circulation and Gas Exchange 896
Excretion and Osmoregulation 899
Nervous System and Sense Organs 900
Reproduction and Development 900
Echinoderm Phylogeny 905
First Echinoderms 905
Modern Echinoderms 908
CHAPTER 27 Phylum Chordata: Cephalochordata and Urochordata 911
Phylum Chordata, Subphylum Cephalochordata: The Lancelets 913
The Cephalochordate Body Plan 913
Body Wall, Support, and Locomotion 913
Feeding and Digestion 915
Circulation, Gas Exchange, and Excretion 915
Nervous System and Sense Organs 916
Reproduction and Development 916
Phylum Chordata, Subphylum Urochordata: The Tunicates 917
The Tunicate Body Plan 920
Body Wall, Support, and Locomotion 925
Feeding and Digestion 925
Circulation, Gas Exchange, and Excretion 927
Nervous System and Sense Organs 927
Reproduction and Development 927
Chordate Phylogeny 931
CHAPTER 28 Perspectives on Invertebrate Phylogeny 935
Illustration Credits IC-1
Selected References SR-1
Index I-1
